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Vol. 35 No 4
BULLETIN
OF THE
TORREY BOTANICAL CLUB
APRIL, 1908
Hepaticae of Puerto Rico
IX. BRACHIOLEJEUNEA, PTYCHOCOLEUS, ARCHlLEJEUNEA, LEUCOLE-
JEUNEA, AND ANOPLOLEJEUNEA
Alexander William Evans
(with plates 6-8)
BRACHIOLEJEUNEA
The genus Brachial ejeunea includes between twenty and thirty
recognized species. About half of these are American, the others
being found in eastern Asia or among the islands of the Pacific.
No species has as yet been reported from Africa. Although essen-
tially tropical in its distribution, the genus reaches its northern
limits in Florida and japan and its southern in Australia and Pata-
gonia. Since the first species described by Spruce is B. laxifolia
(Tayl.) SchifTn., this may be considered the type of the genus. It
was originally described from specimens collected by Jameson in
Ecuador but is now known also from Bolivia.
With scarcely an exception the species of BracJiiohjamca are
found on trees or on rotten logs. In many cases they grow mixed
with other Lejeuneae or with Frullaniae but they sometimes form
extensive mats by themselves. The plants are usually more or
less pigmented and often appear very dark brown or nearly black,
with little or no indication of glossiness. The prostrate stems
cling closely to the substratum by means of numerous rhizoids,
which take their origin from rudimentary discs at the bases of the
underleaves. The branching is at first irregularly pinnate, but the
female plants after flowering usually exhibit a false dichotomy
very much as in Marchcsinia.
The leaves are imbricated and sometimes densely so. In dry
[The Bulletin for March, 1908 (35 : 97-154, A- /-J)» «"* is sued 20 Ap 1908.]
155
156 Evans: Hepaticae of Puerto Rico
plants the lobes are suberect and wrapped around the stem, but as
soon as they become moistened they spread widely (plate 6, fig-
ure i) and assume a more or less squarrose position. The change
in appearance is very similar to what is found in the genus Mastt-
gotejeunea. The lobes are approximately ovate in outline and vary
at the apex from rounded to apiculate or acute. The margin is
usually entire, but, in certain species, tends to be slightly serrulate
in the neighborhood of a female inflorescence. The lobule is
relatively large and consists of two distinct portions approximately
equal in size {figures I, 2). One of these occupies the carinal
region and takes part in the formation of a conspicuous water-sac,
usually about half as long as the lobe. The other is bounded by
the free margin and is closely appressed to the lobe except at the
apical sinus where an opening into the sac is to be observed. The
appressed portion of the free margin bears a series of teeth from
three to ten in number. These teeth vary greatly in size and in
form (figures 7-9), but in the majority of cases each tooth is sev-
eral cells long and so strongly curved toward the lobe that it can-
not be straightened out even by pressure on the cover-glass (fig-
ure 1 2). The outermost tooth, which represents the apex of the
lobule, varies from a single projecting cell to a broad and rounded
projection (figures 10, 11). The hyaline papilla is borne at the
proximal base of this tooth but is displaced to the inner surface of
the lobule, usually appearing two or three cells from the margin.
Beyond the apical tooth the long and shallow sinus extends, form-
ing a very acute angle with the outer part of the keel. The latter
is more or less arched near the base but tends to be incurved where
it meets the postical margin of the lobe. The leaf-cells have thm
walls but distinct trigones (figure 6). These are usually triangu-
lar in outline, two of the sides being convex and one concave.
The thin places between the trigones are relatively wide and rarely
develop intermediate thickenings.
The undivided underleaves are rotund to reniform in outline
and are attached by an arched line (figure i). In certain species
they are distinctly a uricu I ate at the base (figure 1 4); in others
they arc rounded or even cuneate. The margins are sometimes
plane and sometimes more or less re volute ; they vary from entire
to irregularly sinuate but are never distinctly toothed.
Evans : Hepaticae of Puerto Rico ' 157
In the majority of the species two types of branching are to be
observed, very much as in the genus Bryopteris and in certain
other genera of the Lejeuneae Holostipae* In at least one species,
however, all of the branches seem to conform to the Lcjeunea type,
being borne behind leaves with lobules- The Fmllania type of
branching, in which the subtending leaves fail to develop lobules,
is largely restricted to robust vegetative axes. In a branch of this
character the subtending leaf is partly inserted on the main axis
and partly on the branch, the postical base being slightly revolute
(figures 2, 13). The first underleaf is usually distinctly bilobed
and is displaced in such a way that the branch seems to arise from its
axil. It embraces the base of the branch and partially enwraps
the postical base of the subtending leaf. The first leaf is of small
size and complicate-bilobed, but the lobule is explanate and rounded
at the apex. The succeeding leaves and underl eaves are normal
in appearance. The Lejewiea type of branching, even where it
does not occur in the vegetative portion of a plant, is almost
invariably associated with subfloral innovations. An exception to
this condition, however, is found in the remarkable B. sandvi-
censis (Gottsche) Evans, of eastern Asia and the Hawaiian Islands.
In this species the bract behind which a subfloral innovation arises
is wholly destitute of a lobule and is partially inserted on the inno-
vation itself.
The inflorescence in Brackiolejcunea seems to be fairly constant
for a given species and may be dioicous, autoicous, or paroicous.
The female branch varies greatly in length but is usually distinctly
elongated. Subfloral innovations are invariably present and usu-
ally occur in pairs (figures 4, 5) ; in rarer cases only one innova-
tion is developed. The innovations are often floriferous and give rise
to the false dichotomy which is characteristic of the genus. The
bracts are scarcely complicate and the lobe is usually more pointed
than in the leaves. In many species a wing is developed at the
base of the keel (figures 4, 5)- The bracteole is free or nearly so
and varies at the apex from rounded to retuse or shortly bilobed.
The perianth scarcely projects beyond the bracts unless the basal
portion elongates with the development jrf^sp orophyte. It is
^ — -^ ^M M I — ~ ^^^"
J*
* See Evans, Hull. Torrey Club 34 : 559- l 9°$-
t Trans. Conn. Acad. 10: 419. 1900.
158 Evans: Hepaticae of Puerto Rico
approximately obovoid in shape and rounded to truncate at the
apex, with a short beak. It is sometimes terete and sometimes more
or less compressed, but even in the latter case the lateral keels are
never sharp. The postical surface bears from two to five keels, and
the antical surface is either plane or provided with three or four keels
(figures 21, 22). All of the keels are rounded and are never
winged or toothed in any way. In pluriplicate perianths they are
separated from one another by deep grooves, and there is no dif-
ference, except in position, between the lateral keels and the others.
In autoicous and dioicous species the male spikes are long and
often proliferate at the apex. The bracts are imbricated and tend
to be smaller than ordinary leaves, but their lobules are relatively
larger and more strongly inflated. Even here the free margin of
the lobule is usually more or less toothed. The antheridia occur in
pairs, and the bracteoles are found throughout the entire length of
the spike. In paroicous species the bracts are much fewer and are
essentially like normal leaves ; they differ also from the bracts just
described in bearing the antheridia singly.
The present genus is in most respects clearly defined. The
squarrose leaves, the large lobules with teeth along the free mar-
gin, the subfloral innovations usually occurring in pairs, and the
plicate perianth with unarmed keels afford an excellent combina-
tion of generic characters. Certain species, however, as Spruce
has already pointed out, show a relationship with MarcJmima,
while others are even more closely allied to the genus Ptychocoleus
as restricted in the present paper. In most species of Marchesinia
the free margin of the lobule is also denticulate and the sub-
floral innovations occur in pairs, but the leaves are not squarrose
and their lobes are relatively smaller and very different in appear-
ance from those found in Brachiolejeunca. The perianth, more-
over, is distinctly compressed and plane on both antical and postical
surfaces. The characters which separate Brachiolejeunea from
Ptychocoleus will be considered under the latter genus.
Two species of Brachiolejeunca, B. densifolia (Raddi) comb,
nov. jndif. corticalis (Lehm. & Lindenb.) Schiffn., have been re-
f„^ l^t T C,e * 1S Commonl y known as B. bicohr (Nees) Schiff,,., in spite of the
.nnliH f """•"*' demi f° lia Ra <Wi and Jungtrmanma ticolor Nees have long been
cohered S ynony ms . Trevisan restored Raddi's specific name, but has not been fol-
Evans: Hepaticae of Puerto Rico 159
ported from the West Indies. Neither of these has yet been found
in Puerto Rico, but a third species occurs in recent collections from
the island. It is apparently undescribed and may be characterized
as follows :
Brachiolejeunea insularis sp, nov.
Dull-yellowish or brownish-green, sometimes almost black,
scattered or growing in depressed mats : stems 0.25 mm. in diam-
eter, sparingly pinnate, the branches obliquely to widely spread-
ing, similar to the stem but with somewhat smaller leaves, never
microphyllous : leaves closely imbricated, the lobes slightly falcate,
ovate, 1.2-1,7 mm. long, 0.85-1.2 mm. wide, rounded to subcordate
at the base and rounded or very obtuse at the apex, margin entire,
strongly outwardly curved from the antical base to the apex; lobule
ovate-triangular in outline, 0,85-1 mm. long, 0,35-0.45 mm. wide,
the inflated portion forming a conical water-sac about half as long
as the lobe, keel nearly straight from a more or less arched base,
usually forming a continuous line with the postical margin of lobe,
free margin rounded to cordate at the base, usually bearing from
eight to ten more or less distinct teeth, those normally developed
two or three cells lone, one or two cells wide at the base and curved
inward toward the lobe, apical tooth very variable ; cells of lobe
more or less convex, averaging 14 [t at the margin, 28 x 22 fit in the
middle and 35 x 28/^ at the base, intermediate thickenings infre-
quent, oval : underleaves loosely imbricated, plane, broadly orbic-
ular, 0.6-0.7 mm - I° ll g» o<75~0-85 mm. wide, distinctly auriculate
at the base with crenulate auricles, margin otherwise entire or ir-
regularly sinuate, apex broad, truncate to retuse : inflorescence
paroicous : 9 inflorescence borne on a long branch and innovating
on both sides ; bracts erect-spreading, indistinctly complicate and
unequally bifid, the lobe ovate to oblong, 1 .4 mm. long, 0.85 mm.
wide, rounded to obtusely pointed at the apex, margin more or less
sinuate and crispate but not toothed, lobule adnate to lobe for
greater part of its length, ovate to oblong, 0.85 mm. long, 0.35
mm- wide, apex mostly acute, rarely blunt, margin mostly entire but
rarely with a tooth near the apex, wing broad, approximately semi-
circular, entire, usually adnate for its whole length ; bracteole
lowed by subsequent authors. The synonymy of the species is as follows : Fruilanoides
dtmifoha Raddi, Mem. Soc. liai. Modena Fis, 19 : 3 8 - l82 3 J a0: P L 2 < /■ 5-
1829, Jun^trmannia bicohr Nees, in Martius, Fl, Bras. I 1 : 349. 1833, Lejtunta
bicohr Nees, in Montagne, Flor- Boliv. 66; d'Orbigny, Voy, dans 1'Amir, Merid. 7 1 .
*839- Phragmicoma bicohr Nees, in G. L, & N. Syn. Hep. 294. 1845. Ptycko-
toltus dcmifolius Trevis. Mem. 1st. Lomb, III. 4: 405. 1877. Lejtunea {Brachio-
Lejtunta} bicolor Spruce, Hep. Amaz. et. And. 131. 1 884. Brachioitjtunea
*wfor Schiffh. in Engler & Prantl, Nat. Pflanzenfam. i 3 \ 128. 1895.
160 Evans: Hepaticae of Puerto Rico
oblong to obovate, 1.2 mm. long, 0.85 mm. wide, plane or nearly so,
margin entire or irregularly sinuate, apex broad, truncate or sub-
retuse ; perianth slightly exserted beyond the bracts, oblong- obo-
void, 1.5 mm. long, 0.85 mm. wide, rounded to truncate at the apex,
slightly or not at all compressed, mostly ten-keeled, the keels ex-
tending to below the middle, rounded and separated by deep
grooves : c? bracts in one or two pairs below the involucre, essen-
tially like the ordinary leaves : mature sporophyte not seen.
(Plate 6.)
On trees and logs. Near Mayaguez, Heller {4.4.63a), Near
Cayey, Evans (p/). Mount Morales, Utuado, Howe {465). The
writer's specimens from near Cayey may be designated the type.
The species has also been collected in Cuba, Wright, Undenvood &
Earle, and in Jamaica, Underwood \ Evans.
B. insularis is closely related to B. densifolia, and the two
species have been more or less confused. B. dcnsifolia was origi-
nally collected in Brazil, where it seems to be abundant and widely
distributed. Its range extends also along the chain of the Andes
from Colombia to Bolivia. In North America it has been recorded
from Mexico by Gottsche and from the island of St. Vincent by
Spruce. It resembles B. insularis in general habit, in its auriculate
underleaves and in its pluriplicate perianth. It is markedly distinct,
however, in its dioicous inflorescence, the male inflorescences form-
ing long spikes with closely crowded bracts. It differs also in its
greater size, in its more sharply pointed leaves, in the fewer and
smaller teeth along the free margin of the lobule and in its revo-
lute underleaves. Even in B. densifolia the leaves are sometimes
rounded as in B. insularis, but this is an exceptional condition and
is usually associated with incomplete development.
B. corticalis, which is surely to be expected in Puerto Rico, is
considerably smaller than B. insularis, and the teeth along the
free margins of the lobules are fewer and simpler, each tooth con-
sisting usually of a single projecting cell. The species is further
characterized by its dioicous inflorescence, by the more distinct
bracteal lobules, ligulate-Ianceolate in outline and rounded at the
apex, and by the more strongly compressed perianth with fewer
and more irregular keels. Another close ally is B. chinantlana
(Gottsche) Schiftn.,* at present known from Mexico and Ecuador.
* Hedwigia 33 : 180. //. 9 .f. J2 - 40 . ,8 94 . The spec i es i s based on Phragmi
coma buotor var. thinantlana Gottsche, Mex. Leverm. 172. 1863.
Evans: Hepaticae of Puerto Rico 161
This species agrees with B. insularis in its inflorescence, but the
lobes of its leaves and bracts are sharply pointed. The bracteole
is also said to be emarginate or even bifid, but the figure of
Schiffner does not show this condition clearly.
PTYCHOCOLEUS
The history of Trevisan's genus Ptychocoleus has already been
noted by the writer in another connection.* It is based on
Phragmicoma, § Ptychanihoides of the Synopsis Hepaticarum, and
most of its species would be included in the genera Acrolejeunea,
Brachiolejeunea or Mastigolejeunca, as these are at present under-
stood. The first species listed by Trevisan is P. aulacophorus
(Mont.) Trevis. This was originally described by Montagne.f
under the name Phragmicoma aulacophora, from specimens col-
lected in the Mangareva or Gambier Islands, of Spencer Gulf,
South Australia, but is now known to have a wide range extend-
ing into both Asia and Africa. Ptychocoleus aulacophorus would
naturally be selected as the type of the genus. This species, how-
ever, is placed by Stephani in Acrolejeunea, and it therefore seems
justifiable, in restoring the name Ptychocoleus, to apply it to the
genus Acrolejeunea as defined by Spruce and by Schiffner.
In this restricted sense Ptychocoleus would include between
forty and fifty recognized species, all of which are tropical in their
distribution. It attains its best development in southeastern Asia
and the islands of the Pacific but is also well represented in Africa.
In America five species are at present known, three of which have
been reported from the West Indies. Only one species, however,
P. poly car pus (Nees) Trevis., has been collected in Puerto Rico.
As in the preceding genus, practically all the species of Ptycho-
coleus grow on trees or on rotten logs.
The genus is characterized by an elongated female branch
without innovations and by a plicate perianth with smooth keels
(plate 7, figure 2), the number varying from four to ten in dif-
ferent species. The absence of subfloral innovations will at once
distinguish it from Brachiolejeunea, but the two genera agree so
* Bull. Torrey Club 34 : 543- l 9 oS - „ . - .,
t Ann. Sci. Nat. Bot. II. ig : 259. 1S43. Voy. au FOle Sud, Bot. I i 210.pl,
/y./ /. 1845.
1^2 Evans : Hepaticae of Puerto Rico
closely in the characters derived from the lobes of the leaves,
from the underleaves, and from the bracts and perianths that it
would be superfluous to describe these organs in detail for Pty-
chocoleus. There are also no essential differences in color, in
general habit, or in cell-structure. The lobules of the leaves, how-
ever, are much more variable in Ptychocoteus than in Brachiolejeunea,
especially with respect to the number of teeth on the free margin.
Certain species develop as many as four or five teeth, others only
two (figure 5), while in still others the margin is quite entire
except for the apical tooth. In the position of the hyaline papilla
the genus agrees with Brachiolgauiea, except for the fact that it is
sometimes so much displaced that it is difficult to determine
whether it is distal or proximal to the apical tooth, while on the
other hand it may arise directly from a marginal cell. The position,
however, is usually constant for a given species. The branching,
so far as observed, always conforms to the Lejeunea type.
The genus shows but little relationship with other Lejeuneae
which lack subfloral innovations. In Lophokjeunea the keels of
the perianth are winged and variously toothed or laciniate, the
lobules are built up on a different plan, and the local thickenings
in the cell-walls are different. In Caudalejeunea the perianth is
trigonous, its keels are normally winged and toothed, the plants
are different in habit, and the leaf-cells show very numerous inter-
mediate thickenings. In Bryopteris the female branch is much
shorter, the perianth is again trigonous, the leaves and under-
leaves are sharply toothed, and the whole habit, general appear-
ance, and cell -structure are different.
Ptvchocoleus polycarpus (Nees) Trevis.
Jungcrmannia polycarpa Nees, in Martius, Fl. Bras V • 350.
1833.
Phragmicoma polycarpa Nees, in G. L. & N. Syn. Hep. 295. 1845.
Lejatuea domingensis Tayl. Lond. Jour. Bot. 5 : 389. 1846.
Ptychocoleus polycarpus Trevis. Mem. 1st. Lomb. III. 4: 405.
1877.
Lijainea (Aero -Lejeunea) domingensis and polycarpa Spruce, Hep.
Amaz. et And. 116. 1884.
>jettnea polycarpa Schiffn. in Engler & p rant l ( Nat. Pflanzen-
fam. i 3 : 128. ' 1895.
Evans : Hepaticae of Puerto Rico 163
Olive-green, not glossy, scattered or growing in depressed
mats: stems 0.17 mm. in diameter, irregularly pinnate, the
branches widely spreading, occasionally microphyllous : leaves
densely imbricated, the lobe suberect and convolute when dry,
widely spreading and squarrose when moist, more or less convex,
broadly ovate, slightly falcate, 1-1.2 mm. long, 0.85-0.95 mm.
wide, arching partially or wholly across axis, rounded at base,
rounded to very obtuse at the apex, margin entire, strongly out-
wardly curved from the antical base to the apex ; lobule ovate,
0.6 mm. long, 0.35 mm. wide, the inflated portion occupying a
little more than the carinal half, keel arched, forming an obtuse or
rounded angle with the postical margin of lobe, free margin
straight or a little curved, often bearing a tooth one or two cells
long in the vicinity of the apical tooth, the latter mostly two cells
long and one and two cells wide at the base, slightly outwardly
curved, hyaline papilla borne on a marginal cell but slightly dis-
placed to the inner surface of the lobule ; cells of lobe plane or
nearly so, averaging l$p at the margin, 2} x 1$ ft in the middle
and 35 x 18// at the base, intermediate thickenings occasional,
oval or circular: underleaves loosely imbricated, plane or slightly
revolute at the apex, broadly orbicular-obovate, 0.6 mm. long,
O.75 mm. wide, cuneate toward base and attached by an arched
line, broad and truncate at the apex, margin entire or vaguely
angular-denticulate : inflorescence autoicous or polyoicous : $
inflorescence borne on an ascending and more or less elongated
branch ; bracts widely spreading, complicate with a rounded keel
and shortly and unequally bifid, the lobe ovate, 1.6 mm long, 0.85
mm. wide, acute and apiculate, the apex usually tipped with a row
of two or three cells, margin entire, lobule acuminate, otherwise
similar to the lobe, 1.4 mm. long, 0.4 mm. wide; bracteole free,
oblong or ovate, 1.4 mm. long, 0.85 mm. wide, undivided and
acute or shortly bidentate at the apex with sharp or blunt teeth,
margin otherwise entire ; perianth about one third exserted,
obovoid, 1.5 mm. long, O.85 mm. wide, slightly compressed and
cuneate toward the base, truncate to retuse at the apex, antical face
plane or nearly so, postical face with two confluent rounded keels
often extending beyond the beak : cf inflorescence terminating a
more or less elongated simple branch, not proliferating; bracts
mostly in eight to twelve pairs, shortly and unequally bifid with a
strongly arched keel, lobe obliquely spreading, more or less
squarrose, rounded, lobule suberect, mostly acute; antheridia
borne singly ; bracteoles extending the whole length of the spike,
similar to the underleaves but more often revolute at the apex :
capsule about 0.5 mm. in diameter, walls of inner layer of cells
thickened on the inside with a few large circular or oval pits ;
164 Evans : Hepatic ak of Puerto Rico
spores green, spherical or ellipsoidal, 35 /1 in diameter, minutely
verruculose and with circles of indistinct radiating ridges ; elaters
about 10 « in diameter, sometimes bispiral in the middle. (Plate
7, figures 1— 1 1.)
On trees. Between Cayey and Caguas, Howe (1411 p- P->
1414). Type locality, Brazil, Martins ; since found by numerous
collectors. Known also from Mexico, Liebman, and from the fol-
lowing islands of the West Indies: Cuba, Wright; Santo Do-
mingo, Pcrsoon (the type locality of Lejeuttea domingensis); Ja-
maica, Harris, Evans.
The species is not well described in the Synopsis Hepaticarum,
but some of its most significant characters have since been empha-
sized by Schiftner.* Considerable confusion was introduced by
Taylor into the synonymy of the plant by the publication of Le-
jeitnea domingensis. This species was based on specimens which
he at first called L. linguaefolia, and material thus named was
deposited in the Hooker herbarium. Subsequently he renamed
the species L. domingensis and transferred the name L. linguaefolia
to an entirely distinct plant collected by Richard on the island of
St. Thomas. Both species were soon afterwards published in the
same paper. Many years later, Spruce, basing his opinion on the
specimens in the Hooker herbarium rather than on Taylor's de-
scriptions, referred both L. domingensis and L. linguaefolia to his
subgenus Acro-Lejennea, apparently considering them distinct from
each other and also from Ptychocoleus poly carp us. Through the
study of authentic material in the Lindenberg herbarium, Stephani t
was able to reduce both species to synonymy, L. domingensis
proving to be identical with Ptychocoleus polycarpus and the true
L. linguaefolia from St. Thomas proving to be the same as Bra-
chiolcjcunca cortkalis. Specimens in the Mitten herbarium, now in
the possession of the New York Botanical Garden, amply confirm
the conclusions of Stephani.
The keels of the perianth in P. polycarpus scarcely extend
below the middle and are exceedingly variable even on a single
plant According to Schiffner, five or six keels are present, while
Spru ce gives the n umber as four or five. These discrepancies are
*Hedwigia 33: 181. 1894. See also Gottsche, Abhandl. nuturw. Verein.
Bremen 7 r 350, 35^ 1882.
fHedwigiaaS: 166, 167. 1889 ; 39: 22. 1890.
Evans ; Hepaticae of Pukrto Rico 165
doubtless due to differences of interpretation. When the antical
surface is deeply grooved it produces the effect of two rounded
keels, making the total number five or six, according to whether
the confluent postical keels are counted as one or two* When the
antical face is plane the number of keels would naturally be esti-
mated as four.
The two other species of Ptychocolats which have been reported
from the West Indies are P. torulosus (Lehm. & Lindenb.) Trevis.
and Lejatnea {Acrolejeuncd) atroviridis Spruce,* both of which are
listed by Spruce from the island of St. Vincent, where they were
collected by Elliott. The second of these species is known from
no other locality, but P. torulosus is known also from Guiana and
Brazil. In L. atroviridis the trigones of the leaves are absent or
minute and the keels of the perianth are more or less roughened,
so that it is possible that it ought to be referred to some other
genus. P. torulosus, however, is a typical representative of Ptyeho-
cohus. It may be at once distinguished from P. polycarpus by the
blunt lobes and lobules of its bracts, by its truncate bracteole and
by its seven- to nine-keeled perianth. It is distinct also m the
lobules of its leaves, which bear from three to eight teeth along the
free margin instead of one or two.
ARCHILEJEUNEA
According to Schiffner, the genus ArchiLjatnea contained thirty
species in 1895, and perhaps a dozen new species have been pro-
posed since he made his estimate. If, however, A. fordloidcs
(Spruce) SchifTn., the first species described by Spruce, be selected
as the type of the genus, certain species have been referred to it
which can hardly be considered congeneric with this type species.
This is the case, for example, with A. pseudocucullata Steph. {Lejat-
nea holostipa Spruce), which the writer has already made the type of
the genus Cyrtolejeunca^ It is true of A. xanthocarpa (Lehm. &
Lindenb.) Steph. and its immediate allies, for the reception of which
the genus Uucolejeunea Evans has recently been proposed. It is
also true of A. conferta (Meissn.) Schiffn., which belongs to the
genus Anoplolcjeunea and is even synonymous with i ts type species.
- ■■ - ^m ^
^'" '■ ■ ■ 1 ■ - — ta*»_^
* Jour. Linn. Soc. BoL 30 : 335. /A 20. / j-10- 1894.
t Bull. Torrey Club 30 : 55 2 « l 9°3*
166 Evans : Hepaticae of Puerto Rico
If these aberrant forms are removed, the species which are left
will fully agree with the characters assigned to the genus 'by Spruce
and by Schiffner. In the majority of cases the plants are robust
and show a reddish or brownish pigmentation. A prostrate and
irregularly branched caudex is first developed, which clings closely
to the substratum (plate 8, figure 2). From this, secondary
stems soon arise which are more or less free from the substratum
and rarely form rhizoids ; they branch irregularly, many of the
branches being subfloral innovations (figure i). Although the
branches frequently bear smaller leaves than the stem, they are
never truly microphyllous.
The leaves are imbricated but not densely so, and the lobes
spread obliquely or widely from the axis, being more or less fal-
cate. They vary in outline from suborbicular to ligulate, the
apices are rounded and the margins are entire. The lobules, in
their most typical form, are rhomboidal in outline, the keel being
straight or nearly so. In some cases about half of the lobule is
appressed to the lobe, restricting the water-sac to a narrow linear
space along the keel. In other cases the appressed portion is
much narrower (figure 2) and may even be absent altogether, the
free margin meeting the lobe at a right angle. Under these cir-
cumstances the water-sac is relatively larger and the entire lobule
may enter into its formation. The free margin is more or less
curved and terminates in an apical tooth, which varies from suba-
cute to long-acuminate ; the sinus beyond is long but usually
shallow. The hyaline papilla is at the proximal base of the
apical tooth and may be either marginal or slightly displaced to
the inner surface of the lobule. Except for the apical tooth the
margin is commonly entire. In certain species, however, a second
tooth is sometimes developed, proximal in position to the papilla
(figure 4). In some cases the secondary stems fail to develop
normal lobules (figure i), and it becomes necessary to search for
them on the prostrate caudex. The cells of the lobe have firm
and pigmented walls and are usually a little convex. The trigones
are large and conspicuous, and intermediate thickenings are fre-
quent. The trigones are sometimes triangular but it is more
usual for them to be triradiate with rounded rays (figure 3).
The underleaves vary from orbicular to reniform. They are
Evans: Hepaticae of Puerto Rico 167
broad and rounded at the apex, entire along the margin, and vari-
able at the base, being sometimes cuneate and sometimes rounded
or subcordate. The line of insertion is slightly arched and there
is sometimes a rudimentary radicelliferous disc at the base- Ex-
cept on the caudex, however, rhizoids are very scantily de-
veloped.
In the majority of cases the inflorescence is dioicous, but a few
autoicous species have been described. The female inflorescence
is borne on a secondary stem or one of its leading branches and
usually innovates on only one side. In a few species, however,
two subfloral innovations are occasionally developed. An inno-
vation spreads obliquely and frequently forms a second arche-
gonium after bearing only one or two pairs of foliage leaves.
When this procedure is repeated several times in succession, a
cymose flower-cluster is the result, in which the flowers seem to
be borne along the upper side of a floral axis. In rare cases an
innovation is terminated by an antheridial spike. The bracts are
unequally bifid and sharply complicate but are apparently never
winged along the keel. The lobes spread obliquely and are rela-
tively narrower than in the leaves, often showing a tendency to be
sharp-pointed at the apex. The lobule is also narrow and varies at
the apex from rounded to acute, The bracteole is free and ovate to
obovate in outline, the apex showing all variations in different species
from rounded or retuse to bidentate or bifid. In both bracts and
bracteoles the margins are commonly entire. The perianth is
oblong to obovate in outline, with a rounded or truncate apex and
a short beak. It is more or less compressed with sharp lateral
keels (figure 8). The antical surface commonly bears a short
and low keel in the upper part, while the postical surface bears
two sharp and confluent keels extending to below the middle.
Both lateral and postical keels usually develop narrow and inter-
rupted wings which are either irregularly sinuous or angular-
dentate along the margin. In other respects the surface of the
perianth is smooth.
The male inflorescence is large and conspicuous, terminating a
secondary stem or one of its leading branches ; in many cases, how-
ler, it proliferates at the apex. The diandrous bracts are loosely
imbricated and unequally bifid, both lobe and lobule being either
168 Evans: Hepaticae of Puerto Rico
rounded or very obtuse at the apex. The bracteoles extend the
whole length of the spike and are much like the underleaves.
Several genera of the Lejeuneae Holostipae are more or less
closely allied to Archilejeunea. Certain species of Brachiolejeunea
and Ptychocoleus, for example, develop a five-keeled perianth which
is built up on a very similar plan. In Brachiolejeunea, however,
the lobule is differently constructed and shows a larger number of
marginal teeth, while in Ptychocoleus no subfloral innovations are
present. Both of these genera, moreover, are composed of pros-
trate species which show no distinction between caudex and
secondary stems, and which are further characterized by the smooth
and wingless keels of their perianths. In Mastigolejeunea the dis-
tinction between caudex and secondary stems is well marked but
the perianth is sharply trigonous with smooth keels. Spruce
divided his subgenus Archi-Lejeunea into two sections : Monotro-
pella, in which the keels of the perianth are rough and the sub-
floral innovations usually occur singly ; and Dibrachiella, in which
the keels are smooth and the innovations are sometimes borne in
pairs. Schiffher accepts both of these sections, making them sub-
genera of his genus Archilejeunea. Whether the slight differences
just noted will ever be deemed sufficient to separate the groups
generically is doubtful, since they share so many characters in
common. It must be admitted, however, that Dibrachiella shows
an especially close relationship to Brachiolejeunea and Ptycho-
coleus.
In its restricted sense Archilejeunea is almost exclusively trop-
ical in its distribution. A number of species have been described
from Africa and from the islands of the Pacific, but the highest
development of the genus is attained in South America, where
about half of the known representatives have been collected. At
the present time no species are definitely known from Asia and
only two species, both belonging to the subgenus Dibrachiella,
have been reported from the West Indies. One of these is A.
Auberiana (Mont.)Steph., originally described from Cuba, and the
other is A. Cruegeri (Lindenb.) Schiffn., originally described from
Trinidad. The distribution of these two species beyond the islands
where they were first discovered is still imperfectly known.
Stephani, to be sure, has reported A. Auberiana from Paramaribo
Evans : Hepaticae of Puerto Rico 169
and Para, * and Spruce has also reported A. Cruegeri from the
latter locality, f Stephani's report is based on two specimens in the
Lindenberg herbarium, one doubtfully referred to Lejeunea unci-
loba Lindenb. and the other representing a portion of the original
material of L. cydostipa Tayl. The writer has examined both of
these specimens and would refer the first to L. (Arc/ii-Lejeum-a)
florentissima Spruce rather than to A. Anberiana ; % the second is
unfortunately so fragmentary that positive determination is hardly
possible. With regard to A. Cruegeri the type material itself is
scanty and poorly developed, making it difficult to obtain an ade-
quate idea of the species. In view of these facts confirmatory
evidence as to the occurrence of A. Anberiana and A. Cruegeri in
South America is much to be desired. Although Archilcjeunea
has not yet been reported from Puerto Rico, a single species, also
belonging to the subgenus Dibrachiella, has been found on the
island, namely :
Archilejeunea viridissima (Lindenb.)
Lejeunea viridissima Lindenb. in G. L. & N. byn. Hep. 320.
1845.
Lejeunea {Archi- Lejeunea) viridissima Steph. Hedwigia 29: 17.
1890,
Yellowish- or brownish-green, becoming darker with age,
neither glossy nor glaucous, growing in depressed mats : caudex
and secondary stems about O.14 mm. in diameter, the latter simple
or sparingly subdivided, often with poorly developed lobules:
leaves imbricated, the lobe widely spreading, slightly falcate, broadly
ovate, 0.75 mm. long, 0.6 mm. wide, convex, and sometimes revo-
lute at the broad and rounded apex and along the postical margin,
antical margin straight or a little rounded at the base, then strongly
outwardly curved to the apex ; lobule (when well developed) ovate,
O.35 mm. long, 0.25 mm. wide, -the water-sac broad at the base
and abruptly narrowed in the outer part, free margin sometimes
revolute at the base, sometimes appressed to the lobe from base
to apex, rounded, normally bearing two teeth in the outer part,
the outer or apical a little longer than the other and often variously
curved, the inner varying from acute to obtuse an d sometimes
Hedwigia 29 : I5> 2I * 1 890.
t Hep. Atnaz. et And* 97. 1884,
% See Torreya 7 : 228. 1908.
170 Evans : Hepaticae of Puerto Rico
absent altogether, papilla marginal at base of apical tooth, sinus
shallow, keel slightly arched, forming a very obtuse angle with the
postical margin of the lobe ; cells of lobe plane or a little convex,
averaging 12 fi at the margin, 24 x 18/i in the middle and 28 x
18 /i at the base, trigones large, triradiate with rounded ends,
separated from the oblong to circular intermediate thickenings by
narrow pits : underleaves distant to loosely imbricated, appressed
on the caudex but more or less squarrose on the secondary stems
and sometimes revolute at the apex and along the sides, broadly
orbicular-obovate, averaging 0.35 mm. in length and 0.4 mm. in
width, cuneate at the base, rounded to subretuse at the apex: in-
florescence polyoicous : 9 inflorescence terminating a secondary
stem or one of its leading branches, innovating on one side or
rarely on both, the innovations short and sterile or soon again
floriferous, rarely terminating in an antheridial spike ; bracts erect-
spreading, deeply and unequally bifid, complicate, the lobe oblong-
obovate, 0.95 mm. long, 0.5 mm. wide, more or less convex and
often revolute along the postical margin and at the apex, lobule
ovate to ligulate, 0.4 mm. long, o. 1 5 mm. wide, rounded to subacute
at the apex, keel very short ; bracteole obovate, averaging o. 7 5 X 0.4
mm., rounded to retuse at the apex; perianth about half exserted,
oblong-obovate in outline, cuneate toward the base, 1 .4 mm. long,
0.75 mm. wide, antical keel low and broad, lateral and postical keels
high and sharp, usually bearing narrow wings from one to three cells
wide and variously sinuate orsubcrenulate on the margin : c? inflo-
rescence terminating a short branch, apparently not proliferating ;
bracts loosely imbricated, mostly in from three to six pairs, une-
qually bifid, the lobe oblong to rounded at the apex, lobule similar
but shorter and narrower, keel strongly arched ; antheridia in
pairs ; bracteoles similar to the underleaves but smaller : capsule
about 0.4 mm. in diameter, yellowish-brown ; spores greenish,
irregular in form, about 16/* in short diameter, minutely verrucu-
lose ; elaters 9,0 in diameter. (Plate 8, figures 1-8.)
On a log. El Yunque, Evans (140). Type locality: Caracas,
Venezuela (herb. Hampe). The species has also been found on
the island of Great Bahama, Britton & Milhpmigh. The type
material preserved in the Lindenberg herbarium agrees closely
with the Puerto Rico and Bahamian specimens.
Archdejeitnea viridissima is closely related to A. Auberiana.
Through the kindness of M. Paul Hariot the writer has been sup-
plied with a portion of the original material of the latter species
from the Montagne herbarium. It shows an autoicous inflores-
cence and agrees with A. viridissima in size, habit, and color.
Evans: Hepaticae of Puerto Rico 171
The lobules in both species are normally bidentate, and the
perianths are five-carinate with smooth or nearly smooth keels.
The lobule in A. Auberiana, however, is subrectangular in outline
rather than ovate, the keel being nearly straight, and the marginal
teeth tend to be larger and more pronounced than in A. viridis-
sima. The leaf-cells also offer certain points of distinction,
although they average about the same in size. In A. Aubcriana
the local thickenings of the walls are rarely confluent, the trigones
are mostly in the form of equilateral triangles with straight sides,
and the intermediate thickenings are oval or circular in outline.
In A. viridissima, on the other hand, the thickenings are more
frequently confluent, the trigones show a strong tendency to be
triradiate, with rounded and often constricted rays, and the inter-
mediate thickenings show a similar tendency to be rectangular,
with rounded ends. A. Aiiberiana is further characterized by the
lobules of its perichaetial bracts ; these are much smaller than in
A. viridissima and are usually adnate to the lobes throughout
their entire length. Although the differences which have just
been enumerated are slight, they seem to be constant.
Another allied species is Lejennea florcntissima Spruce, which
the writer would keep distinct from A. Auberiana. This species
is more robust than A. viridissima, and the lobes of the leaves
are more falcate ; the keel of the lobule is parallel with the free
margin, but both are more or less curved; the trigones in the leaf-
cells are similar to those in A, Aubcriana but are better developed,
the sides of the triangles being convex rather than straight. The
subfloral innovations in L. florcntissima are usually soon floriferous
and the perianths are thus crowded together in a cymose cluster;
in A. viridissima the arrangement of the perianths is much more
irregular.
LEUCOLEJEUNEA
The characters of the genus Leucolejeunta, recently segregated
by the writer from Archil ejeune a, have already been so fully dis-
cussed * that it hardly seems necessary to enumerate them here.
Five species are definitely known at the present time, the generic
type being L. clypeata (Schwein.) Evans, a widely distributed
species in the eastern United States. The only representative of
*See Torreya 7 ; 225-229, 1908.
172 Evans : Hepaticae of Puerto Rico
the genus which has been found in Puerto Rico is Z, xanthocarpa,
although L. unciloha (Lindenb.) Evans is surely to be expected
on the island,
Leucolejeunea xanthocarpa (Lehm. & Lindenb.) Evans
Jungermannia xanthocarpa Lehm. & Lindenb. in Lehmann, Pug,
Plant. 5:8. 1832,
Lejeunea xanthocarpa Lehm. & Lindenb.; G. L. & N. Syn. Hep.
330, 1845.
Lejeunea {Archi-Lejettned) xanthocarpa Pears, Christiana Vidensk.-
Selsk, Forh. 1887 9 : 4. pL i.f. 14.-24,
Archil ejettnea xanthocarpa Schiffn. Conspect. Hepat Archip. Indici
316. 1898.
Leucolejeunea xanthocarpa Evans, Torreya 7: 229. 1908.
Pale glaucous-green, varying to bright-green and becoming
yellowish or brownish upon drying, growing in depressed mats :
stems 0.15 mm. in diameter, the branches obliquely to widely
spreading, with smaller leaves than the stem but not microphyl-
lous, sterile branches mostly simple or sparingly subdivided : leaves
imbricated and usually densely so, the lobe (when flattened out)
orbicular-ovate, 1 mm. long, 0,85 mm- wide, convex and strongly
revolute at the broad and rounded apex and along the whole pos-
tical margin, antical margin straight or slightly rounded at base,
then outwardly curved to the apex ; lobule narrowly ovoid, 0.4
mm. long t 0.2 mm. wide, gradually contracted in the outer half,
the water-sac opening directly into the revolute portion of the lobe,
keel straight or slightly arched continuous with revolute portion or
slightly indented at junction, apex (not visible without flattening
or dissection) varying from bluntly pointed to acuminate, in the
latter case tipped with a row of from two to four cells, sinus straight
or slightly iunulate, hyaline papilla mostly three to six cells from
the apex ; cells of lobe plane or slightly convex, averaging 14/***
the margin, 20 p in the middle, and 28 x 20 (i at the base : under-
leaves imbricated, plane (or a little convex from below), reniform,
0.5 mm. long, 0,7 mm, wide, rounded to cordate at the base,
rounded to vaguely retuse at the apex : 9 inflorescence borne on
a more or less abbreviated branch, sometimes with only one pair
of leaves below the involucre, innovating on one side ; bracts
erect-spreading, the lobe falcate, convex, revolute at the rounded
apex and along the postical margin, oblong-obovate, 1 mm, long,
0.6 mm. wide, lobule oblong, 0,4 mm. long, 0.2 mm. wide, rounded
to subacute at the apex, keel sometimes narrowly winged; brac-
teole broadly obovate, 0.95 mm. long, 075 mm. wide, slightly
Evans : Hepaticae of Puerto Rico 1 73
retuse at the apex ; perianth almost hidden by the involucre, I mm.
long, 0.7 mm. wide, narrowed toward the base, rounded to slightly
retuse at the apex, beak 0.1-0.15 mm. long, keel roughened by
projecting cells but destitute of wings or distinct teeth : o 71 in-
florescence as described under the genus. (Plate 7, figures
12-23.)
On trees. Near Cayey, Evans (76). The species is also
known from the following islands of the West Indies: New
Providence, Bahamas, A. E. Wight, E. G. Britton ; Jamaica, Un-
derwood, Evans; Guadeloupe, Hn snot, Buss ; Dominica, Elliott;
Martinique, Duss. On the American continent its range extends
from Mexico to Brazil (the type locality). There are also records
from Java and from Fernando Po, Mount Kilimanjaro and Cape
Colony in Africa. A fragment of the type specimen in the Lin-
denberg herbarium, collected by Beyrich in 1823 at New Freiburg,
Brazil, agrees closely with the specimens here described.
Apparently the closest allies of L. xantkocarpa tare L. unciloba,
which has already been mentioned, and L. conchifolia Evans.*
Both of these species have convex lobes and long teeth at the
apices of the lobules. In L. unciloba, however, the apex of the
lobe is only slightly revolute, and the apical tooth of the lobule is
easily visible without dissection. In L. conchifolia the plants are
less robust than in L. xantkocarpa, the lobes of the leaves spread
more obliquely, the lobules are relatively larger, and their apical
teeth are more like those of L. unciloba, being readily demonstrated
without dissection. Z. xantkocarpa also bears a strong superficial
resemblance to Pycnolejeuuca Sckivamckei (Steph.) Schiffii., which
is of course at once distinguished by its bifid underleaves.
ANOPLOLEJEUNEA
The genus Anoplolcjeunca, as defined by Schifiner, contains the
single species A. herpestica (Spruce) Schiffii. Apparently nothing
has been written about this species except the rather brief descrip-
tions of Spruce and Schiffner. The writer finds, however, upon
examining a portion of the type material of A. herpestica^ kindly
^^descriptions and figures of these two species see Evans, Mem. Torrey Club
8: 125-129. pi. 16. /. it-zo ; pi f?. / '"V- ^02. L. unciloba is there called
ArehiUjennea Stllowiana Steph., a species which the writer now considers synonymous
with the much older Ltjeunea unciloba Lindenb. See also Torreya 7 I 229. 190S.
174 Evans : Hepatic ae of Puerto Rlco
sent by Mr. M. B. Slater, that it is quite indistinguishable from the
widely distributed Lejeunea conferta Meissn. Although placed by
recent writers in Archilejeunea, L. conferta cannot be considered
congeneric with the typical members of this genus, and there seem
to be excellent reasons for maintaining Anoplolejeuma as distinct
Spruce compares it with his subgenus Plaiy-Lejetinea f but it is
evidently much more closely related to the genus Leitcolejeunea*
The plants grow in depressed mats and are either pale- or dark-
green in color with neither glossiness nor pigmentation. The pros-
trate stem is copiously and irregularly branched, some of the
branches being similar to the stem while others are ascending and
distinctly microphyllous (plate 8, figure 9). The normal leaves
are imbricated, the convex lobe spreading widely and showing a
rounded apex and entire margin. The leaf-cells are plane or
nearly so, with small trigones and occasional intermediate thicken-
ings (figure 1 6), and ocelli are sometimes present at the base of
the lobe (figure 17).
The lobule when normally developed is more complicated than
in most of the other genera of the Lejeuneae* It is ovate-oblong
in outline and forms a strongly inflated water-sac with an arched
and convex keel (figure 9). From a relatively short base it is
abruptly dilated and becomes suddenly contracted in the outer
part. The dilated portion, bounded by the rounded and entire
free margin, forms a volute with about one and a half turns (fig-
ure 14). This comes into contact with the lobe, thus enclosing
the water-sac, but the margin itself, together with a considerable
extent of the lobule, lies free within the sac and can be distin-
guished by careful focusing from below. The lobule develops no
apical tooth, the free margin being directly continuous with the
sinus; just at the junction of the two, however, a cell may be
observed which is a little larger than its neighbors and which
doubtless represents the terminal cell of the lobule, homologous
with the apical tooth of other Lejeuncae (figure 18). At the
beginning of its course the sinus is also strongly revolute but
straightens out abruptly and passes into the postical margin of the
lobe at the end of the keel. The hyaline papilla is marginal and
arises from the terminal cell, usually at its junction with the first
cell of the sinus ; it may therefore be regarded as distal in posi-
Evans : Hepaticae of Puerto Rico 175
tion. Unfortunately, the lobule is often poorly developed and
fails to show some of the peculiarities which have just been
described.
The underleaves are distant and smaller than is usual among
the Holostipae. They are obovate and entire, the apex varying
from truncate to slightly retuse. When rhizoids are developed
they usually spring from a distinct basal radicelliferous disc.
The inflorescence is dioicous, and the male and female plants
usually occur in separate mats. The female branch varies greatly
in length and nearly always give rise to one or two subfloral inno-
vations. In rare cases, however, the branch remains perfectly
simple. The perichaetial bracts are a little larger than the leaves
and their lobes are relatively narrower ; the lobules are rounded
to subacute and the sharp keels are narrowly winged. Thebracteole
is free and mostly obovate-oblong in outline, the apex varying from
rounded to slightly retuse (figures 19, 21 ). The perianth is obo-
void and normally shows five sharp keels (figure 23) and a short
but distinct beak. The male inflorescence occupies a short branch
or is terminal on a more or less elongated branch. The bracts
are imbricated and diandrous, and the bracteoles are limited to the
base of the spike.
Anoplolejeunea conferta (Meissn.)
Jungermannia conferta Meissn.; Sprengel in Linnaeus, Syst. Veg.
ed. 16. 4: 325. 1827.
Lejeunea subaurita Nees & Mont. Ann. Sci. Nat. Bot. II. 5 : 6a
1836.
Lejeunea Miquclii Lehm. & Lindenb. in Lehmann, Pug. Plant. 7:
15- 1 838.
Lejeunea proteoides Lehm. & Lindenb, /. c. 7 : 19. 1838.
Lejeunea conferta Meissn. ; G. L. & N. Syn. Hep. 333. 1845-
Lejeunea involutiloba Mont Ann. Sci. Nat. Bot. IV. 5 : 3 50- 1856.
Lejeunea (Anopio- Lejeunea) herpestica Spruce, Hep. Amaz. et And.
129. 1884.
Lejeunea {Platylejeunea) conferta Steph. Hedwigia 27 : 284.
1888.
Lejeunea (Arc hi- Lejeunea) conferta Steph. /. c. 29 : 21. 1890.
Anoplolejeunea herpestica Schiffn. in Engler & Prantl, Nat. Pflan-
zenfam. I 3 : 131. 1895.
176 Evans : Hepaticae of Puerto Rico
Archilejeiuiea confcrta Schiffh. Conspect. Hepat Archip, Indici
315. 1898.
Plants becoming yellowish or brownish upon drying: stems
0.17 mm. in diameter, branches obliquely to widely spreading:
leaves imbricated, the lobe widely spreading, often revolute at the
apex, falcate, broadly ovate, 1 mm. long, 0.85 mm. wide when of
maximum size, often considerably smaller, arching across or a little
beyond the axis, antical margin straight or slightly incurved near
the base, then strongly outwardly curved to the apex, postical
margin straight or somewhat curved, not revolute, forming an
angle of 90 degrees or more with the strongly arched keel ; lobule
0.35 mm. long, 0.2 mm. wide ; cells of lobe averaging 17 /i at the
margin, 25 ft in the middle and 40 X 25 //at the base, trigones
triangular with concave sides, intermediate thickenings sometimes
confluent with the trigones ; ocelli (when present) mostly one to
three, situated near the base of the lobe, measuring about 45 X
32 fi: underleaves broadly orbicular-obovate, 0.35 mm. long,
O.35-0.4 mm. wide, plane, cuneate and short-decurrent at the base :
perichaetial bracts erect-spreading, the lobe obovate from a nar-
row base, 1,2 mm. long, 0.7 mm. wide, convex and revolute at the
apex, rounded to obtuse at the apex, lobule ovate-oblong, 0.75
mm. long, 0.25 mm. wide; bracteole 0.4-0.75 mm. long, 0.25-
0.4 mm. wide; perianth about two-fifths exserted at maturity,
1-1,3 mm - long, 0.7-0.75 mm. in diameter, apex variable, usually
truncate or subretuse but sometimes rounded or even subacute,
keels variable, sometimes low and almost obsolete, sometimes dis-
tinctly dilated in the upper part, when well developed showing a
marginal row of slightly projecting cells with strongly thickened
walls, postical surface of perianth sometimes developing one or two
low and supplementary keels or folds : cT inflorescence apparently
never proliferating ; bracts mostly in from two to seven pairs,
closely imbricated, inflated, shortly bifid, the lobe strongly convex,
rounded at the apex, antical margin straight or a little incurved,
keel strongly arched, lobule obtuse to acute at the apex ; bracte-
oles similar to the underleaves but smaller: mature sporophyte
not seen. (Plate 8, figures 9-23.)
On trees. Puerto Rico, without definite locality, SinUnis
{rod). The writer has seen no specimens of A. confcrta from
Puerto Rico but has collected it abundantly on the island of
Jamaica, where it occurs at elevations of from 1,000 to 2,000 m.
The species was originally collected in Peru but is now also known
from Brazil, Bolivia, Venezuela, Trinidad, Colombia, and Mexico.
Whether it occurs outside of tropical America is somewhat doubt-
Evans: Hepaticae of Puerto Rico 177
ful. The Synopsis Hepaticarum cites it from the islands of Luzon
and Java, and SchifTner also quotes the latter locality. Both of
these records, however, are based on old collections.
The microphyllous branches of A. conferta (figure 9) are very
characteristic, although they are sometimes scantily developed and
may even be absent altogether. The lobes of their leaves are orbic-
ular-ovate in outline and average about O.35 mm. in length ; they
are suberect or spread obliquely from the axis instead of spread-
ing widely as on normal stems and branches. The lobules are
nearly spherical and measure only o. 1 5 mm. in diameter, in other
respects agreeing with those on typical leaves. The underleaves,
except for their small size, present no distinctive peculiarities.
Under certain circumstances the lobules are poorly developed even
on the stems and leading branches (ficure i 2) and thus give rise to
forms which are not always easy to recognize. Usually a careful
examination will reveal a few lobules of normal structure scattered
among the others, and these will aid in the determination of such
anomalous specimens. The characters upon which the varieties
Miquelii and Liebmaniana of the Synopsis are based are apparently
drawn from imperfectly formed lobules, and the propriety of
attempting to maintain them is doubtful.
In the specimens from Jamaica the lobes of the leaves are
usually distinctly ocellate at the base except on poorly developed
individuals. In all the South American specimens, however, which
the writer has been able to examine, the ocelli are apparently ab-
sent. If this difference should prove to be constant it might
necessitate the separation of the West Indian plant as a distinct
species. Unfortunately the South American specimens were all of
old collections, and the detection of ocelli in plants of this charac-
ter is sometimes very uncertain. Under the circumstances it seems
best to include the Jamaican specimens with the others, a course
which is advocated also by Stephani.
A. conferta bears a rather strong superficial resemblance to
Leucolejeunea clypeata, the two species agreeing in color and in
many of the characters derived from leaf-cells, underleaves, pen-
chaetial bracts, and perianths. They differ strikingly however, m
the structure of the lobule, and L. clypeaia is also distinct m its
inflorescence, which is normally autoicous, and in its lack ot micro-
178 Evans: Hepaticae of Puerto Rico
phyllous branches. There is little danger of confusing A. conferta
with any other species of Leucolejeunea, in spite of the close re-
lationship of the two genera.
The present paper concludes the discussion of the Lejenmae
Holostipae which are known to occur in Puerto Rico. Of the
genera recognized by SchifTner the following four are apparently
unrepresented on the island : Pty chant 1ms % Thysananthus, Peltole-
jeunea and Dicranolejeunea. The last of these genera is abundant
on Jamaica at rather high altitudes and may therefore be expected
to have a wider distribution in the West Indies. There is little
probability, however, that any of the others will be found there,
although each is represented in South America by one or more
species. Ptychanthus and Thysananthus, in fact, are essentially
paleotropic in their distribution and attain their highest develop-
ment in the East Indies and the neighboring parts of Asia.
Yale University.
Evans; Hepaticae of Puerto Rico 179
Explanation of plates ft-£
As in the previous papers of this series the figures were drawn by the writer and
prepared for publication by Miss Hyatt,
Plate 6
Brachi&hjeunea insularis Evans. Part of stem with branch, postical view,
X 15- 2. Part of stem with base of branch, postical view, X ! S« 3- ^ eaf » antical
view, X l S- 4? S* Perianths with involucres, the innovations dissected away,
postical view, X 25. 6. Cells from middle of lobe, X 26 5- 7~9- Fr ^e margins of
lobules, X 45* IO > ll - Apices of lobules, X 20 °- I2 * Marginal teeth of lobule,
X 200. 13. Stem-leaf at base of branch, X *5- H- Basal auricle of underleaf,
X 200. 15-17* Bracts and bracteole from one involucre, X f 5« 18-20. Bracts and
bracteole from another involucre, X *5* 21, 22. Transverse sections of perianths,
X35- Fig- 5 was drawn from specimen collected by Heller {44&3 a ) 5 Fi S s * *• 9»
II, 18-20, from specimens collected by Howe (46J) ; the remaining figures from the
type specimen*
Plate 7
Ptychccoleus polycarpm (Nees) Trevis. r. Part of stem with base of a branch,
postical view, X f 5" 2 ' A P ex of f ema]e branch with perianth, postical view, X J 5-
3, Part of stem, antical view, X<5* 4- Cells from middle of lobe, X 2 65« 5- Margin
of lobule, the apex on right, X 2°°- 6 ^ S * Bracts and b ^cteole from the same invo-
lucre, X J 5- 9- Subfloral leaf below involucre, X l S< l °- Bracteole from another
specimen, X l 5* **« Transverse section of perianth, X 2 5- The Sg ures weTe a11
drawn from specimens collected by Howe (1411* 1414}*
Leucolejeunea xanthwarpa (Lehm. & Undent) Evans. 12. Part of sterile
stem, postical view, X *5- *3- part o f P ]ant witl1 perianth, postical view, V 25* 14*
Part of stem, antical view, X *5- *5- Leaf with lobule flattened out, X 15 l6 * Cel1s
from middle of lobe, X265* 17. Cells from antical margin of lobe, X 200, 1 8, 19-
Apices of lobules, X 200," 20-22. Bracts and bracteole from the same involucre,
X25, 23. Transverse section of perianth, X 35* The figures were all drawn from
specimens collected by the writer (76).
Plate 8
Architejtunea viridhiima (Lindenb.) Evans. I. Apex of female stem with peri-
anth, postical view, X 2 5- 2 > part of caudex, postical view, X 2 5- 3- Cells from
middle of lobe, X 26 5- 4- Margin of lobule, the apex on right, X 2< *>- 5i 6 - Bract
and bracteole from the same involucre, X 2 5- 7* Bract from another involucre, X 2 5-
8. Transverse section of perianth, X 2 5- The figures were all drawn from specimens
collected by the writer (140)*
AnoptoUjtunta conftrta ( Meissn. ) Evans. 9- P» rt of sterile stem with ^'W 1 "
lous branches, postical view, X 25. 10. Female branch with perianth, postical view,
X25. II. Apex of male branch, postical view, X »* I2 - Branch Wlth P °° t } Y de "
veloped lobules, postical view, X 25. *3- Lobule, postical view, X 45- »4- Longi-
tudinal section through lobe and lobule, X 45- *5- Mouth of lobule, diagrammatic. lb.
Cells from middle of lobe, X 265. U- Cells from base of lobe, enclosing an ocellus.
X200. 18. Margin of lobule, apical end on left, X200. IQ. Perianth with invo-
lucre, postical view, X 35- 20, at. Bract and bracteole from a single involucre,
X 25- 22. Perianth, postical view, X 25. ** Transverse section of penantb, X 35-
The figures were all drawn from Jamaican specimens collected by the writer (jj, 272,
424)'
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